Microhabitat Use, Seasonal Activity and Diet of the Snake-Eyed Skink (Ablepharus Kitaibelii ﬁtzingeri)Incomparison with Sympatric Lacertids in Hungary

Biologia, Bratislava, 62/4: 482—487, 2007 Section Zoology DOI: 10.2478/s11756-007-0092-6

Microhabitat use, seasonal activity and diet of the snake-eyed skink (Ablepharus kitaibelii ﬁtzingeri)incomparison with sympatric lacertids in Hungary

Gábor Herczeg1,TiborKovács1,ZoltánKorsós2 & János Tor¨ ok¨ 1

1Behavioural Ecology Group, Department of Systematic Zoology and Ecology, E¨otv¨os Loránd University, Pázmány Péter sétány 1/C, H-1117 Budapest, Hungary; e-mail: [email protected] 2Department of Zoology, Hungarian Natural History Museum, Baross u. 13,H-1088 Budapest, Hungary

Abstract: Microhabitat selection and seasonal activity of the snake-eyed skink, Ablephaus kitaibelii fitzingeri,arecompared to the two lacertid lizards (Lacerta viridis and Podarcis muralis) that co-occur in many of its habitats. The food composition of A. k. fitzingeri is also described. Significant differences in microhabitat selection and seasonal activity among the three species were found. The snake-eyed skink was associated with open grasslands, and with a low level of scrub, bare soil and rock cover. The microhabitat preference of L. viridis was quite similar to that of the skink, but with a higher preference for scrub. P. muralis occurred in places with greater rock and bare soil cover, and more scrub than A. k. fitzingeri. Activity of the snake-eyed skink decreased dramatically in summer, probably because of the reduced thermal inertia originating from the extremely small size of this species, but its seasonal activity overlapped with those of the lacertids. Stomach content analysis of the snake-eyed skink suggests that it is a generalist predator of small, mainly flightless arthropod prey. Competition with juvenile lacertids and predation by adult L. viridis are conceivable for the snake-eyed skink. Key words: Ablepharus kitaibelii fitzingeri; diet composition; Lacerta viridis; microhabitat use; Podarcis muralis;seasonal activity

Introduction rent distribution and possible threatening factors of A. k. fitzingeri in Hungary was published recently (Herczeg The snake-eyed skink, Ablepharus kitaibelii Bibron et et al. 2004). Its general ecology is virtually unknown Bory, 1833 is the only representative of the genus in except from some anecdotal observations, and even its Europe and also the northernmost European represen- current demographic status is based on inadequate cen- tative of the family Scincidae. Despite the unique status sus data, and most local populations are subject to of the snake-eyed skink in the European herpetofauna, continued human disturbance (Korsós 1994; Herczeg & almost all aspects of its biology have remained unstud- Korsós 2003). Before any sound conservation plan can ied (Gasc et al. 1997; Pasuljevič 1965, 1976), proba- be developed, a basic knowledge of its general ecology bly because of its secretive habits. Ablepharus kitaibelii is required. fitzingeri Mertens, 1952, which occupies the northern Community structure and resource utilisation / range of the species’ distribution area, occurs mainly partition patterns of lizards have been an attrac- in Hungary, with sporadic records in southern Slovakia tive topic for ecologists in the last half century (e.g., and northern Serbia (Mertens 1952; Fuhn 1969; Gruber Schoener 1968; Pianka 1973; Vitt et al. 1981, 2000). 1981; Ljubisavljevic et al. 2002). From this work it was suggested that space, time and In Europe, the species is listed in the Bern Con- diet are perhaps the most important limiting factors vention (Council of Europe 1994), Appendix 2 and in where segregation in lizard assemblages could be de- the European Union Habitat Directive (European Com- tected (e.g., Schoener 1968; Pianka 1973; Toft 1985; mission 1992), Annex IV, as a species in need of strict Vrcibradic & Rocha 1996). The tools developed in these protection. A. k. fitzingeri is strictly protected in Hun- studies are widely used to answer conservation ques- gary and is listed in the Hungarian Red Data Book tions (e.g., Sartorius et al. 1999; Vega et al. 2000) or as a potentially endangered taxon (Rakonczay 1989). for describing the ecological requirements of endangered However, most previous studies have dealt only with species (e.g., Martín & Salvador 1995). In this paper, its morphology and distribution (e.g., Fitzinger 1829; we present the first report of microhabitat use, sea- Lendl 1899; Fejérváry 1912, 1917, 1925; Bolkay 1914; sonal activity and diet of the snake-eyed skink. With Méhely 1918; Fejérváry-Lángh 1943; Szunyoghy 1954; respect to microhabitat use and seasonal activity, we Dely 1978; Solti & Varga 1988). A summary of the cur- compared A. k. fitzingeri with the sympatric Lacerta

c 2007 Institute of Zoology, Slovak Academy of Sciences Ecological notes on Ablepharus kitaibelii fitzingeri 483 viridis Laurenti, 1768 and Podarcis muralis Laurenti, the sample sizes were too small for biologically meaningful 1768, the lacertid lizards that co-occur with the snake- interpretations. Taxonomic diet composition was summa- eyed skink at most of its Hungarian sites (Herczeg & rized as the proportion of prey items from a given taxon Korsós 2003). Our goal was to provide some ecological in the total number of prey items (n%) and the proportion data on the snake-eyed skink, which conservationists of lizard individuals eating a prey taxon (F), after James (1991) and Maragou et al. (1996). can use for the conservation management of this reptile endemism of the Carpathian Basin. Statistical analyses We used principal components analysis to reduce the six Material and methods original environmental variables to a smaller number of or- thogonal principal components (PCs). We used data only from spring and autumn as A. k. fitzingeri was almost com- Study organisms and sampling pletely absent in summer (n = 2). Three PCs were extracted The northernmost subspecies (A. k. fitzingeri)ofthesnake- from the six original variables according to Kaiser’s criteria. eyed skink (A. kitaibelii) is one of the smallest lizards in To gather biologically interpretable PCs we rotated the ini- Europe, with a snout to vent length (SVL) of 20–55 mm tial factor solutions by the Varimax procedure. We tested (juveniles ca. 22–30 mm) and body mass (BM) of 0.15–1.5 for differences in the PC scores between species using Anal- g. In Hungary, it occurs in different habitats with respect ysis of Variance (ANOVA) followed by the LSD test. Due to substrate: sandstone, limestone, dolomite, andesite, gab- to the low number of observed A. k. fitzingeri individuals, bro, basalt, or even pure sand (Herczeg et al. 2004; for more we did not test for seasonal differences. We compared the details of the species see Gruber 1981). The common wall seasonal activities of the species using χ2 tests. lizard (P. muralis) is a rock-dwelling, insectivorous helio- We calculated Shannon diversity indices and compared therm species and has SVL of 25–65 mm (juveniles ca. 25–35 them with Hutcheson t-tests (Hutcheson 1970). To compare mm) and BM of 0.7–6.5 g, and is widespread in Europe (for the diet composition between the spring and autumn sam- details see Gruschwitz & Böhme 1986). The green lizard (L. ples, we calculated Proportional Similarity indices. To ex- viridis) is one of the largest lizards in Europe with SVL of plore the relationship between F and n%, we used the Spear- 30–120 mm (juveniles ca. 30–45 mm) and BM of 0.9–25 g. It man rank correlation test. is a ground-dwelling, insectivorous heliotherm lizard and is Because our purpose was to study interspecific differ- widespread in Europe (for details see Nettmann & Rykena ences, we randomly assigned individuals between sex, age 1984). and size categories. All statistics were computed using STA- Fieldwork was carried out in the Sas Hill Nature Re- TISTICA 7.0 for WINDOWS (StatSoft Inc., Tulsa, Okla- serve, a dolomite hill (maximum elevation 259 m a.s.l.) homa, 1994) software. within the area of Budapest (47◦30 N; 19◦51 E). The habitat offered various microhabitat types, such as closed or opened dolomite grasslands, solitary or aggregated scrub, Results forest patches and rocky outcrops. We sampled lizards over six, five and six days in spring, The three PCs together accounted for 75% of the total summer and autumn, respectively, from 12 March to 12 variance (Table 1). According to Norman & Streiner November in 2002 to investigate microhabitat use and sea- (2000), the critical value for the minimum acceptable sonal activity. Sampling days were once every second week factor loading was 0.362 in our case. The first PC was within each season and we chose mostly clear, sunny days. negatively correlated with cover and height of arbo- Sampling was always by the first author from 07.00 to 19.00 real plants and positively with the cover of herbaceous on each day. GH moved slowly through the area and if a lizard was detected, species and microhabitat variables plants (Table 1). Thus PC1 describes a gradient from (cover of herbaceous plants, cover of arboreal/mainly scrub/ grasslands to scrub and forest patches. The second PC plants, rock cover and percentage of bare soil were estimated was positively correlated with the cover and height of to the nearest 5%; height of herbaceous plants and arboreal herbaceous plants and negatively with rock cover (Ta- plants were measured (to the nearest 5 cm) and recorded ble 1), likewise describing a gradient from grassy areas within 2 m of the location where the lizard was first seen. A to rocky outcrops. The third PC was positively corre- constant effort was made to sample each habitat type. Al- lated with the cover of bare soil and negatively with the together 255 lizards (46 A. k. fitzingeri, 113 L. viridis,and cover of herbaceous plants (Table 1), i.e., it represents 96 P. muralis) were recorded. Some additional individuals a gradient from open, patchy grassland to thick, closed were observed without full data recording. We used the latter data only in the seasonal activity analyses. We did not grassland. mark (or capture) the lizards, but, due to the large study The ANOVAs on the PC scores revealed significant area and the high lizard density, we assumed that the rate effects of the factor lizard species for both PCs (Fig. 1; of repeated observations of the same individuals is low. PC1: F2,202 = 3.84; P = 0.02; PC2: F2,202 = 32.78; P We used the standard stomach-flushing methods (Legler < 0.001; PC3: F2,202 = 3.38; P = 0.04). In PC1 A. k. & Sullivan 1979; James 1990) using tubes with diameter fitzingeri differed from both lacertids (LSD tests; all P ranging from 0.7 to 1.2 mm (with respect to the size of < 0.04) while the latter did not differ from each other the given lizard specimen) for the diet analysis. Forty-one (LSD test: P = 0.51). In PC2 and PC3 P. muralis dif- individuals of A. k. fitzingeri were stomach-flushed, 20 in fered from A. k. fitzingeri and L. viridis (LSD tests: spring, and 21 in autumn. Lizards for stomach-flushing were < collected on days other than those used for recording micro- all P 0.03), while the latter did not differ from each habitat use and activity data. None of the sampled individ- other (LSD tests: all P > 0.58). The seasonal activities uals suffered injuries or died because of the sampling. We differed among the species in all pairwise comparisons, 2 made an effort to sample the diet of the lacertids also, but even after a Bonferroni adjustment (χ16 > 68.03, P < 484 G. Herczeg et al.

Table 1. Correlations of microhabitat variables with the ﬁrst three PC scores after Varimax rotation for A. k. ﬁtzingeri, L. viridis and P. muralis using data from spring and autumn.

PC1 PC2 PC3

Ground cover Herbaceous plants 0.481 0.600 −0.625 Arboreal plants −0.886 −0.112 −0.039 Bare soil 0.087 0.196 0.942 Rock cover 0.030 −0.914 0.057 Height Herbaceous plants −0.037 0.482 0.215 Arboreal plants −0.799 0.151 0.036 Eigenvalue 1.667 1.502 1.332 Cumulative % of variance 0.278 0.528 0.750

Fig. 2. Seasonal activity of A. k. ﬁtzingeri, L. viridis and P. muralis.

with a low proportion of bushes and trees, bare soil or rocks. When compared to the skink, P. muralis pre- ferred less closed grasslands with a higher proportion of scrub, and was obviously associated with the rocky outcrops, while L. viridis occupied similar grasslands to the skink but with a higher proportion of bushes and trees. We note that our results are based on microhabitat variables recorded in an area within 2 m of the observed lizards, thus to an extent, the species co-occur. The phylogenentic history of the species stud- Fig. 1. Microhabitat selection of A. k. fitzingeri (Akf), L. viridis ied adds an important component to the understanding (Lv) and P. muralis (Pm). Mean PC scores ± SE provided. For of lizard communities (e.g., Vitt et al. 2003); further, thedescriptionofthePCsseetextandTable1. lizard morphology and habitat-use are related as a result of adaptation (Vanhooydonck & Van Damme 1999; Vanhooydonck et al. 2000). A. k. fitzingeri is a typical 0.001; Fig. 2). Prey diversity of A. k. fitzingeri did not skink with its slim, cylindrical body and reduced legs. differ between spring and autumn (Hutcheson t-test: P It avoids rocky places, as its locomotion is less effective > 0.9). We found a high level of overlap between the on non-horizontal rock surfaces due to its morphology, diet composition of A. k. fitzingeri in spring and au- and it also needs loose soil for digging. P. muralis is tumn (Proportional Similarity index: 0.67). F and n% morphologically (and evolutionarily) adapted to a rock- were strongly correlated (rS = 0.95, n = 13, P < 0.001). dwelling life, thus its association with rocky microhabi- The most frequent prey taxa (in order of n%) were Ho- tats with low scrubs is not surprising. Microhabitat use moptera, Araneae, Formicidae and Coleoptera in the of L. viridis is harder to interpret. It appears to move skink’s diet (Table 2). well in both microhabitat types, with adult individuals moving more than 5–10 m during normal escaping Discussion and foraging events in a few seconds. However, its association with scrub can be derived from another be- We found that A. k. fitzingeri occurred in grasslands havioural characteristic. We observed considerable dif- Ecological notes on Ablepharus kitaibelii fitzingeri 485

Table 2. Stomach contents of A. k. ﬁtzingeri. Columns do not necessarily denote equivalent taxonomic levels. Number of prey items, proportion of the total number of prey items (n%), and the proportion of lizard individuals eating the prey taxon (F) are shown. Marked (bold) prey taxa represent altogether more than 75% of the total number of prey items of A. k. ﬁtzingeri in its whole activity season. (L) denotes larvae.

Season Spring Autumn Total

Shannon diversity 2.01 2.06 2.16 Sample size 20 21 41 Prey Nn%FNn%F N n%F

Insecta Homoptera Aphidina 15 21 20 3 4.8 9.5 18 13.3 14.6 Others 17 23.6 45 11 17.5 33.3 28 20.7 39 Heteroptera 1 1.4 5 3 4.8 14.3 4 3 9.8 Coleoptera 8 11.1 15 8 12.7 28.6 16 11.8 21.9 Coleoptera (L) 3 4.2 15 4 6.3 19 7 5.2 17.1 Hymenoptera Formicidae 2 2.8 10 17 27 28.6 19 14.1 19.5 Others11.45– – – 1 0.72.4 Lepidoptera 1 1.4 5 – – – 1 0.7 2.4 Lepidoptera (L) 5 6.9 20 3 4.8 14.3 8 5.9 14.1 Diptera 2 2.8 10 2 3.2 9.6 4 3 9.8 Diptera (L) – – – 1 1.6 4.8 1 0.7 2.4 Arachnida Araneae 14 19.4 60 11 17.5 47.6 25 18.5 58.5 Pseudoscorpiones 3 4.2 10 – – – 3 2.2 4.9

Total 72 100 – 63 100 – 135 100

ferences in the escape tactics between the skink and predation pressure on the skink cannot be disregarded, the lacertids (Herczeg & Korsós 2003), and these differ- as in the stomach contents of eight adult L. viridis,we ences might influence their microhabitat use. The lac- found remnants of two adult A. k. fitzingeri. ertids flee without cover to the nearest refuge, which is Seasonal activity patterns were different among the usually a rock crevice for P. muralis and scrub for L. three species, but the activity of A. k. fitzingeri over- viridis (we found that L. viridis often climbs to scrub lapped to a large extent with the two lacertid species. when chased), thus the latter should prefer grassland The minimal activity of A. k. fitzingeri in summer, the with solitary scrub or the grassland – scrubland edges hottest and driest season, suggests an increased danger (see also Korsós 1984). A. k. fitzingeri needs no special of overheating for this extremely small-bodied species, refuge, as it usually responds to predators (e.g., human due to its small thermal inertia (Herczeg et al. 2007). observer) by immediately hiding within 1 m under grass The bimodality in the activity of L. viridis is a result and leaf litter and probably digging itself in, hence it of the bimodal activity of the juvenile (and thus sim- can occur with no regard to scrub. These results sug- ilarly constrained) members of the species (Herczeg et gest that a common habitat-depressing factor, namely al. 2007). the fragmentation of grasslands by spreading natural In summary, A. k. fitzingeri was associated with or invasive scrub species (Herczeg et al. 2004), would grasslands and avoided continuous rock surfaces and favour L. viridis. dense scrub, and dramatically decreased its activity in A. k. fitzingeri was found to be a generalist preda- summer. Our results support the conclusions of our pre- tor from the high diversity of its prey and the corre- liminary study (Herczeg & Korsós 2003): namely that, lated F and n% values. Prey taxa eaten by A. k. fitzin- although there are differences in the spatial and tem- geri were also important components of the diet of dif- poral distributions of the studied species, the overlaps ferent lacertids as found in previous studies (e.g., Av- among them are considerable. A. k. fitzingeri was found ery 1966; Koponen & Hietakangas 1972; Valakos et al. to be a generalist predator of small, mainly flightless 1997). However, the great importance of the taxon Ho- arthropod prey. Juvenile lacertids, and especially L. moptera for the skink is unusual. In contrast, important viridis (due to the spatial overlap with A. k. fitzin- taxa in the diet of different lacertid species such as Gas- geri), could be its competitors, while adult L. viridis tropoda, Orthoptera and Isopoda (Avery 1966; Kopo- individuals were its predators. In the light of our re- nen & Hietakangas 1972; Diaz & Carrascal 1990; Mel- sults, we suggest that in the conservation management lado & Corti 1993; Rugiero 1994; Valakos et al. 1997) of A. k. fitzingeri, the population interactions within were completely absent from the stomach contents of A. a focal lizard assemblage cannot be disregarded. In k. fitzingeri. We assume that, according to the head size anthropogenically-disturbed habitats, the disturbance – prey size relationship in most generalist lizards (e.g., due to habitat conversion might change the strength of DeMarco et al. 1985; Vitt 2000), juvenile lacertids, ir- the interpopulation interactions in many ways (Vega et respective of species, could compete for food with A. al. 2000; Taylor & Fox 2001) possibly causing declines in k. fitzingeri. In addition, in the case of adult L. viridis, populations of this potentially endangered subspecies. 486 G. Herczeg et al.

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